Marie F. Arndal

Soil and plant community characteristics and dynamics at Zackenberg

Arctic soils hold large amounts of nutrients in the weatherable minerals and the soil organic matter, which slowly decompose. The decomposition processes release nutrients to the plant-available nutrient pool as well as greenhouse gases to the atmosphere. Changes in climatic conditions, for example, changes in the distribution of snow, water balance and the length of the growing season, are likely to affect the complex interactions between plants, abiotic and biotic soil processes as well as the composition of soil micro- and macro-fauna and thereby the overall decomposition rates. These interactions, in turn, will influence soil-plant functioning and vegetation composition in the short as well as in the long term. In this chapter, we report on soils and. plant communities and their distribution patterns in the valley Zackenbergdalen and focus on the detailed investigations within five dominating plant communities. These five communities are located along an ecological gradient in the landscape and are closely related to differences in water availability. They are therefore indirectly formed as a result of the distribution of landforms, redistribution of snow and drainage conditions. Each of the plant communities is closely related to specific nutrient levels and degree of soil development including soil element accumulation and translocation, for example, organic carbon. Results presented here show that different parts of the landscape have responded quite differently to the same overall climate changes the last 10 years and thus, most likely in the future too. Fens represent the wettest sites holding large reactive buried carbon stocks. A warmer climate will cause a permafrost degradation, which most likely will result in anoxic decomposition and increasing methane emissions. However, the net gas emissions at fen sites are sensitive to long-term changes in the water table level. Indeed, increasing maximum active layer depth at fen sites has been recorded together with a decreasing water level at Zackenberg. This is in line with the first signs of increasing extension of grasslands at the expense of fens. In contrast, the most exposed and dry areas have less soil carbon, and decomposition processes are periodically water limited. Here, an increase in air temperatures may increase active layer depth more than at fen sites, but water availability will be critical in determining nutrient cycling and plant production. Field manipulation experiments of increasing temperature, water supply and nutrient addition show that soil-plant interactions are sensitive to these variables. However, additional plant-specific investigations are needed before net effects of climate changes on different landscape and plant communities can be integrated in a landscape context and used to assess the net ecosystem effect of future climate scenarios.

Seasonal Variation in Gross Ecosystem Production, Plant Biomass, and Carbon and Nitrogen Pools in Five High Arctic Vegetation Types

Abstract The Arctic is extremely vulnerable to projected climate change, and global warming may result in major community reorganizations. The aim of this study was a thorough investigation of plant biomass production throughout an entire growing season in five different high arctic vegetation types: Cassiope, Dryas, and Salix heath, grassland, and fen. The main focus was on the gross ecosystem production (GEP), and the biotic and abiotic factors which may influence GEP. Photosynthesis, aboveground biomass, and carbon, nitrogen, and chlorophyll content were measured weekly during nine weeks. There were large differences in seasonal growth and production within and among vegetation types. Mosses contributed considerably to the total C and N pool in grassland, fen, and Salix heath. Fen, which had the highest pool of leaf N, leaf chlorophyll, and moss N, was the most productive vegetation type in terms of GEP, despite the lowest total biomass. Across vegetation types, leaf biomass, leaf N, and moss N pool size influenced GEP. Within most vegetation types GEP correlated with leaf N, in correspondence with the notion that N may limit plant production in many high arctic ecosystems. The timing of the peaks in C and N pools in leaves did not coincide with that in the mosses and in woody tissues. This emphasizes the importance of sampling throughout the growing season, when using field data from the Arctic to estimate plant biomasses and modeling C and N fluxes and pool sizes.

Root growth and N dynamics in response to multi-year experimental warming, summer drought and elevated CO2 in a mixed heathland-grass ecosystem

Ecosystems exposed to elevated CO2 are often found to sequester more atmospheric carbon due to increased plant growth. We exposed a Danish heath ecosystem to elevated CO2, elevated temperature and extended summer drought alone and in all combinations in order to study whether the expected increased growth would be matched by an increase in root nutrient uptake of NH4+-N and NO3- -N. Root growth was significantly increased by elevated CO2. The roots, however, did not fully compensate for the higher growth with a similar increase in nitrogen uptake per unit of root mass. Hence the nitrogen concentration in roots was decreased in elevated CO2, whereas the biomass N pool was unchanged or even increased. The higher net root production in elevated CO2 might be a strategy for the plants to cope with increased nutrient demand leading to a long-term increase in N uptake on a whole-plant basis. Drought reduced grass root biomass and N uptake, especially when combined with warming, but CO2 was the most pronounced main factor effect. Several significant interactions of the treatments were found, which indicates that the responses were nonadditive and that changes to multiple environmental changes cannot be predicted from single-factor responses alone.

Improved UV-B screening capacity does not prevent negative effects of ambient UV irradiance on PSII performance in High Arctic plants. Results from a six year UV exclusion study.

Long-term responses of ambient solar ultraviolet (UV) radiation were investigated on Salix arctica and Vaccinium uliginosum in a High Arctic heath ecosystem in Zackenberg, northeast Greenland. Over a period of six years, UV exclusion was conducted in the growing season by means of filters: 60% UV-B reduction, 90% UV-B+UV-A reduction, UV transparent filter control, and an open control without filter. Plant responses were evaluated using specific leaf area, leaf content of UV-B absorbing compounds and PSII performance parameters derived from chlorophyll-a fluorescence induction curves. Based on the JIP-test, we calculated the total performance index PI(total), which includes the integrating antennae, the PSII reaction center, intersystem electron transport and reduction of PSI end acceptors-dependent parameters. In both species, UV exclusion significantly decreased the content of UV-B-absorbing compounds. Salix increased its specific leaf area, while Vaccinium decreased it. UV exclusion increased the PI(total) in both species during all six years of experimentation. This response was governed by a significantly decreased RC/ABS, a marginally non-significant increased ET(o)/TR(o) and a significantly increased TR(o)/ABS=F(V)/F(M) and RE(o)/ET(o). These results demonstrate the current level of ambient UV-B to decrease PSII performance significantly in these High Arctic plants. It appears that the two plant species both have improved their UV-screening capacity, but through different strategies, although this did not sufficiently prevent negative effects of the ambient UV radiation. We argue the decreased PSII performance to be part of a response decreasing plant carbon uptake. We speculate the negative effects on PSII performance mediated by ambient UV irradiance to be present in years where warming induces early snowmelt, exposing the vegetation to high spring UV-B, and to be present in the future to the degree the ozone layer is not fully recovered.

Solar UV-B effects on PSII performance in Betula nana are influenced by PAR level and reduced by EDU: results of a 3-year experiment in the High Arctic

The long-term and diurnal responses of photosystem II (PSII) performance to near-ambient UV-B radiation were investigated in High Arctic Betula nana. We conducted an UV exclusion experiment with five replicated blocks consisting of open control (no filter), photosynthetic active radiation and UV-B transparent filter control (Teflon), UV-B-absorbing filter (Mylar) and UV-AB-absorbing filter (Lexan). Ethylenediurea (EDU), a chemical normally used to protect plants against ozone injury, was sprayed on the leaves both in the field and in an additional laboratory study to investigate if EDU mitigated the effects of UV-B. Chlorophyll-a fluorescence induction curves were used for analysis of OJIP test parameters. Near-ambient UV-B radiation reduced across season maximum quantum yield (TR(o) /ABS = F(v) /F(m)), approximated number of active PSII reaction center (RC/ABS) and the performance index (PI(ABS)), despite improved leaf screening against UV-B with higher content of UV-B-absorbing compounds and a lower specific leaf area. EDU application counteracted the negative impact of UV-B on TR(o) /ABS, RC/ABS and PI(ABS) . This indicates that the mechanisms behind UV-B and ozone damage share some common features. The midday depression was present in all treatments, but TR(o) /ABS and PI(ABS) were persistently lower in near-ambient UV-B compared to UV-B reduction. The recovery phase was particularly impaired in near-ambient UV-B and interactive effects between treatment × hour raised TR(o) /ABS, RC/ABS and PI(ABS) higher in reduced UV-B compared to near-ambient UV-B. This demonstrates current solar UV-B to reduce the PSII performance both on a daily as well as a seasonal basis in this High Arctic species.

Fine Root Growth and Vertical Distribution in Response to Elevated CO2, Warming and Drought in a Mixed Heathland–Grassland

Belowground plant responses have received much less attention in climate change experiments than aboveground plant responses, thus hampering a holistic understanding of climate change effects on plants and ecosystems. In addition, responses of plant roots to climate change have mostly been studied in single-factor experiments. In a Danish heathland ecosystem, we investigated both individual and combined effects of elevated CO2, warming and drought on fine root length, net production and standing biomass by the use of minirhizotrons, ingrowth cores and soil coring. Warming increased the net root production from ingrowth cores, but decreased fine root number and length in minirhizotrons, whereas there were no significant main effects of drought. Across all treatments and soil depths, CO2 stimulated both the total fine root length (+44%) and the number of roots observed (+39%), with highest relative increase in root length in the deeper soil layers. Our results suggest that under future climate, plants may allocate considerable resources into roots compared to aboveground biomass. Increased carbon (C) allocation to roots may have a great impact on the overall ecosystem C balance and must be considered in modelling of future ecosystem responses to climate change. To provide models with necessary validation data, more studies are needed to investigate if higher C allocation to roots will lead to long-term C storage in more recalcitrant soil C pools or if this potential increase in soil carbon storage may be offset by increased priming activity and turnover rates for soil organic matter.

Long-term and realistic global change manipulations had low impact on diversity of soil biota in temperate heathland

In a dry heathland ecosystem we manipulated temperature (warming), precipitation (drought) and atmospheric concentration of CO2 in a full-factorial experiment in order to investigate changes in below-ground biodiversity as a result of future climate change. We investigated the responses in community diversity of nematodes, enchytraeids, collembolans and oribatid mites at two and eight years of manipulations. We used a structural equation modelling (SEM) approach analyzing the three manipulations, soil moisture and temperature, and seven soil biological and chemical variables. The analysis revealed a persistent and positive effect of elevated CO2 on litter C:N ratio. After two years of treatment, the fungi to bacteria ratio was increased by warming, and the diversities within oribatid mites, collembolans and nematode groups were all affected by elevated CO2 mediated through increased litter C:N ratio. After eight years of treatment, however, the CO2-increased litter C:N ratio did not influence the diversity in any of the four fauna groups. The number of significant correlations between treatments, food source quality, and soil biota diversities was reduced from six to three after two and eight years, respectively. These results suggest a remarkable resilience within the soil biota against global climate change treatments in the long term.

Fast Responses of Root Dynamics to Increased Snow Deposition and Summer Air Temperature in an Arctic Wetland

In wet tundra ecosystems, covering vast areas of the Arctic, the belowground plant biomass exceeds the aboveground, making root dynamics a crucial component of the nutrient cycling and the carbon (C) budget of the Arctic. In response to the projected climatic scenarios for the Arctic, namely increased temperature and changes in precipitation patterns, root dynamics may be altered leading to significant changes in the net ecosystem C budget. Here, we quantify the single and combined effects of 1 year of increased winter snow deposition by snow fences and summer warming by open-top chambers (OTCs) on root dynamics in a wetland at Disko Island (West Greenland). Based on ingrowth bags, snow accumulation decreased root productivity by 42% in the 0–15 cm soil depth compared to ambient conditions. Over the growing season 2014, minirhizotron observations showed that root growth continued until mid-September in all treatments, and it peaked between the end of July and mid-August. During the season, plots exposed to experimental warming showed a significant increase in root number during September (between 39 and 53%) and a 39% increase in root length by the beginning of September. In addition, a significant reduction of root diameter (14%) was observed in plots with increased snow accumulation. Along the soil profile (0–40 cm) summer warming by OTCs significantly increased the total root length (54%), root number (41%) and the root growth in the 20–30 cm soil depth (71%). These results indicate a fast response of this ecosystem to changes in air temperature and precipitation. Hence, on a short-term, summer warming may lead to increased root depth and belowground C allocation, whereas increased winter snow precipitation may reduce root production or favor specific plant species by means of reduced growing season length or increased nutrient cycling. Knowledge on belowground root dynamics is therefore critical to improve the estimation of the C balance of the Arctic.